Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)

The different familial assignations proposed until now for Geobatracbus are compiled. Through the collection of a virtual topotypical series of this monotypic genus, which is endemic of the NW section of the Sierra Nevada de Santa Marta Massif (Departamento del Magdalena, Colombia), a detailed descr...

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Autores:
Ardila Robayo, María Cristina
Tipo de recurso:
Article of journal
Fecha de publicación:
1979
Institución:
Universidad Nacional de Colombia
Repositorio:
Universidad Nacional de Colombia
Idioma:
spa
OAI Identifier:
oai:repositorio.unal.edu.co:unal/44306
Acceso en línea:
https://repositorio.unal.edu.co/handle/unal/44306
http://bdigital.unal.edu.co/34404/
Palabra clave:
Ciencias Naturales
Biología
Plantas
animales
Historia Natural
Paleobotánica
Paleozoología
Ornitología
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
Rights
openAccess
License
Atribución-NoComercial 4.0 Internacional
id UNACIONAL2_17d0913a7e90bf91a5a8bc8eeb3a1a35
oai_identifier_str oai:repositorio.unal.edu.co:unal/44306
network_acronym_str UNACIONAL2
network_name_str Universidad Nacional de Colombia
repository_id_str
dc.title.spa.fl_str_mv Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
title Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
spellingShingle Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
Ciencias Naturales
Biología
Plantas
animales
Historia Natural
Paleobotánica
Paleozoología
Ornitología
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
title_short Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
title_full Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
title_fullStr Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
title_full_unstemmed Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
title_sort Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)
dc.creator.fl_str_mv Ardila Robayo, María Cristina
dc.contributor.author.spa.fl_str_mv Ardila Robayo, María Cristina
dc.subject.proposal.spa.fl_str_mv Ciencias Naturales
Biología
Plantas
animales
Historia Natural
Paleobotánica
Paleozoología
Ornitología
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
topic Ciencias Naturales
Biología
Plantas
animales
Historia Natural
Paleobotánica
Paleozoología
Ornitología
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
Amphibia
Anura
Género Geobatrachus Ruthven
Adenomera
description The different familial assignations proposed until now for Geobatracbus are compiled. Through the collection of a virtual topotypical series of this monotypic genus, which is endemic of the NW section of the Sierra Nevada de Santa Marta Massif (Departamento del Magdalena, Colombia), a detailed description was made.  67 morphological characters were selected according to the criteria used by Lynch and Heyer in their studies on the philogeny and classification of the leptodactylid frogs, and such characters, compled with the chromosome number (2n =20) were analysed, after initial examination of their meaning in order to ascertain the affinities of the genus. Field work carried on in the region showed that its distribution cover the area of humid forest, with frequent mists, at. 1.750-3.000 m., and pointed also that the species tolerates paraclimacic situations (in plantations of Cupressus sp., and Pinus sp.) , and their habits are exclusively terrestrial and semifossorial.  Among the external characters it can be emphasized the high degree of chromatic individual variation, the absence of xeromorphic adaptations and specialized glandular complexes in the skin, which point towards an adaptation towards a hygrophilic habitat, added to the extreme reduction of the first pedial digit. The skeleton characters, studied through dissection and differential staining, show a generalized type without pronounced reduction in the cranial elements, the occurrence of non-pedicellated teeth in the maxillary arch, the absence of either phragmotic or fossorial adaptations, or epicraneal ornamentation exostosis; a generalized hyoid apparatus; 8 presacral vertebrae without fusion or imbrication, and with specialized transverse processes; no expanded sacral diapophyses, fused talus and calcaneus and only two tarsalia. The T-shaped terminal phalanges might suggest a preadaptation towards an arboreal way of life (or, conversely, an arboreal ancestor); also the full firmisternal condition of the pectoral girdle and the striking clavicular reduction which would show an apparent relationship with the Microhylids. The following muscular characters are noteworthy: M. intermandibularis with medial raphe, the type "s" insertion of the M. adductor mandibularis subexternus, the distal tendon of M. semitendinosus passing ventral to the Mm. graciles, the insertion over the knee of the M. adductor longus, and the absence of an accessory head (or tendon) in the M. glutaeus magnus (an unique condition among the Neobatrachia, presumably due to secondary loss).  In contrast with previous wrong assertions, a direct development (without free larval stages) was confirmed, a condition expected a priori since the eggs are megalecythal and laid in reduced numbers in protected places, in accordance with the scare opportunities for aquatic larval development in mountain environments.  Each of the 68 used characters were codified according to their primitive or derived represented stages, following criteria adopted in the recent literature, with the aim of using principles of numeric taxonomy to at temp the reconstruction of the phylogeny. The possibility that Geobatrachus could belong to the Microhyloidea due to its firmisternal condition, was easily discarded, and the absence of intercalary skeletal pieces in the digits, prevent also any possible close relationship with the Centrolenidae, Hylidae and Pseudidae, although there are no characters (except for the absence of an accessory tendon in the M. glutaeus magnus, regarded as a secondary loss) that might exclude the inclusion of Geobatrachus in the Bufonoidea. No particular characters allow the inclusion of the genus either in the Myobatrachidae or the Heleophrynidae.  The absence of Bidder's organ (which is the only diagnostic difference between the Leptodactylidae and the Bufonidae) eliminate the family Bufonidae from any further consideration, a conclusion that is supported also by experimental cladograms. Distinctive features such as the morphology of the pectoral girdle and the presence of dorsal osteoderms, characteristic of the Brachycephalidae (sensu McDIARMID, 1969) also exclude this family from consideration.   The combinational analysis made through cladograms confirms the presumptive derivation of the Phyllobatinae (= Dendrobatidae) from the Elosiini, and leads to the conclusion that this group might be better regarded as a subfamily of the Leptodactylidae than as a family of their own. Although, in particular the firmisternal condition might suggest affinities of Geobatrachus with the Phyllobatinae, neither this genus or Rhinoderma can be referred to this subfamily. The analysis of the characters of Rhinoderma shows rather conclusively that this genus was derived from the immediate anc estor of the Leptodactylinae or from early members of this group, and under such circumstances Rhinoderma can equally be included in the Leptodactylidae as a representative of a monotypic subfamily. Otherwise, the Leptodactylidae should be divided in several groups of familial hierarchy. However, Geobatrachus widely differs from the Rhinodermatinae, and, as its clearly shown by the cladograms, shows its greater affinity with the Eleutherodactylini, although Geobatrachus is rather outstandingly different from the other genera of this tribe. This condition might be due to the in situ evolution of an early stock, isolated since the Tertiary, that has survived in favored conditions due to climatic equability. Several hypothesis based on the available paleogeographic and paleoclimatic evidence are discussed, in conjunction with the possible fundamental dichotomy of the" Alpha" and" Beta" groups of the genus Eleutberodactylus. With the noticeable exceptions of the genera Euparkerella and Holoaden, the Eleutherodactylini (perhaps originally issued in S. E. Brazil) represent a group of closely related genera, although an exhaustive review of Eleutherodactylus might eventually lead to the splitting of this genus, and to conclusions that agreeing essentially with the conclusions here offered, could otherwise provide basis for a better interpretation of the facts here presented, or even to a reduction of the currently recognized genera for this tribe.  Both, the firmisterny, the loss of the accessory tendon of the M. glutaeus magnus, and the caryotipe (with comparatively reduced number of chromosomes and the lack of acrocentrics), suggest that Geobatrachus represents an extreme line of specialization among the Eleutherodactylini. The evidence afforded by the cladistic analysis gives no support to regard the Eleutherodactylini are derived from Adenomera or an immediate common ancestor with this genus, and so the acquisition of the direct development among the Leptodactylidae seens to have occurred independently at least in two phyletic lines.
publishDate 1979
dc.date.issued.spa.fl_str_mv 1979
dc.date.accessioned.spa.fl_str_mv 2019-06-28T13:18:12Z
dc.date.available.spa.fl_str_mv 2019-06-28T13:18:12Z
dc.type.spa.fl_str_mv Artículo de revista
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dc.relation.ispartof.spa.fl_str_mv Universidad Nacional de Colombia Revistas electrónicas UN Caldasia
Caldasia
dc.relation.ispartofseries.none.fl_str_mv Caldasia; Vol. 12, núm. 59 (1979); 383-495 Caldasia; Vol. 12, núm. 59 (1979); 383-495 2357-3759 0366-5232
dc.relation.references.spa.fl_str_mv Ardila Robayo, María Cristina (1979) Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura). Caldasia; Vol. 12, núm. 59 (1979); 383-495 Caldasia; Vol. 12, núm. 59 (1979); 383-495 2357-3759 0366-5232 .
dc.rights.spa.fl_str_mv Derechos reservados - Universidad Nacional de Colombia
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dc.rights.license.spa.fl_str_mv Atribución-NoComercial 4.0 Internacional
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spelling Atribución-NoComercial 4.0 InternacionalDerechos reservados - Universidad Nacional de Colombiahttp://creativecommons.org/licenses/by-nc/4.0/info:eu-repo/semantics/openAccesshttp://purl.org/coar/access_right/c_abf2Ardila Robayo, María Cristina5049fa2a-f1f8-43d9-8d2c-ab7393f99fa53002019-06-28T13:18:12Z2019-06-28T13:18:12Z1979https://repositorio.unal.edu.co/handle/unal/44306http://bdigital.unal.edu.co/34404/The different familial assignations proposed until now for Geobatracbus are compiled. Through the collection of a virtual topotypical series of this monotypic genus, which is endemic of the NW section of the Sierra Nevada de Santa Marta Massif (Departamento del Magdalena, Colombia), a detailed description was made.  67 morphological characters were selected according to the criteria used by Lynch and Heyer in their studies on the philogeny and classification of the leptodactylid frogs, and such characters, compled with the chromosome number (2n =20) were analysed, after initial examination of their meaning in order to ascertain the affinities of the genus. Field work carried on in the region showed that its distribution cover the area of humid forest, with frequent mists, at. 1.750-3.000 m., and pointed also that the species tolerates paraclimacic situations (in plantations of Cupressus sp., and Pinus sp.) , and their habits are exclusively terrestrial and semifossorial.  Among the external characters it can be emphasized the high degree of chromatic individual variation, the absence of xeromorphic adaptations and specialized glandular complexes in the skin, which point towards an adaptation towards a hygrophilic habitat, added to the extreme reduction of the first pedial digit. The skeleton characters, studied through dissection and differential staining, show a generalized type without pronounced reduction in the cranial elements, the occurrence of non-pedicellated teeth in the maxillary arch, the absence of either phragmotic or fossorial adaptations, or epicraneal ornamentation exostosis; a generalized hyoid apparatus; 8 presacral vertebrae without fusion or imbrication, and with specialized transverse processes; no expanded sacral diapophyses, fused talus and calcaneus and only two tarsalia. The T-shaped terminal phalanges might suggest a preadaptation towards an arboreal way of life (or, conversely, an arboreal ancestor); also the full firmisternal condition of the pectoral girdle and the striking clavicular reduction which would show an apparent relationship with the Microhylids. The following muscular characters are noteworthy: M. intermandibularis with medial raphe, the type "s" insertion of the M. adductor mandibularis subexternus, the distal tendon of M. semitendinosus passing ventral to the Mm. graciles, the insertion over the knee of the M. adductor longus, and the absence of an accessory head (or tendon) in the M. glutaeus magnus (an unique condition among the Neobatrachia, presumably due to secondary loss).  In contrast with previous wrong assertions, a direct development (without free larval stages) was confirmed, a condition expected a priori since the eggs are megalecythal and laid in reduced numbers in protected places, in accordance with the scare opportunities for aquatic larval development in mountain environments.  Each of the 68 used characters were codified according to their primitive or derived represented stages, following criteria adopted in the recent literature, with the aim of using principles of numeric taxonomy to at temp the reconstruction of the phylogeny. The possibility that Geobatrachus could belong to the Microhyloidea due to its firmisternal condition, was easily discarded, and the absence of intercalary skeletal pieces in the digits, prevent also any possible close relationship with the Centrolenidae, Hylidae and Pseudidae, although there are no characters (except for the absence of an accessory tendon in the M. glutaeus magnus, regarded as a secondary loss) that might exclude the inclusion of Geobatrachus in the Bufonoidea. No particular characters allow the inclusion of the genus either in the Myobatrachidae or the Heleophrynidae.  The absence of Bidder's organ (which is the only diagnostic difference between the Leptodactylidae and the Bufonidae) eliminate the family Bufonidae from any further consideration, a conclusion that is supported also by experimental cladograms. Distinctive features such as the morphology of the pectoral girdle and the presence of dorsal osteoderms, characteristic of the Brachycephalidae (sensu McDIARMID, 1969) also exclude this family from consideration.   The combinational analysis made through cladograms confirms the presumptive derivation of the Phyllobatinae (= Dendrobatidae) from the Elosiini, and leads to the conclusion that this group might be better regarded as a subfamily of the Leptodactylidae than as a family of their own. Although, in particular the firmisternal condition might suggest affinities of Geobatrachus with the Phyllobatinae, neither this genus or Rhinoderma can be referred to this subfamily. The analysis of the characters of Rhinoderma shows rather conclusively that this genus was derived from the immediate anc estor of the Leptodactylinae or from early members of this group, and under such circumstances Rhinoderma can equally be included in the Leptodactylidae as a representative of a monotypic subfamily. Otherwise, the Leptodactylidae should be divided in several groups of familial hierarchy. However, Geobatrachus widely differs from the Rhinodermatinae, and, as its clearly shown by the cladograms, shows its greater affinity with the Eleutherodactylini, although Geobatrachus is rather outstandingly different from the other genera of this tribe. This condition might be due to the in situ evolution of an early stock, isolated since the Tertiary, that has survived in favored conditions due to climatic equability. Several hypothesis based on the available paleogeographic and paleoclimatic evidence are discussed, in conjunction with the possible fundamental dichotomy of the" Alpha" and" Beta" groups of the genus Eleutberodactylus. With the noticeable exceptions of the genera Euparkerella and Holoaden, the Eleutherodactylini (perhaps originally issued in S. E. Brazil) represent a group of closely related genera, although an exhaustive review of Eleutherodactylus might eventually lead to the splitting of this genus, and to conclusions that agreeing essentially with the conclusions here offered, could otherwise provide basis for a better interpretation of the facts here presented, or even to a reduction of the currently recognized genera for this tribe.  Both, the firmisterny, the loss of the accessory tendon of the M. glutaeus magnus, and the caryotipe (with comparatively reduced number of chromosomes and the lack of acrocentrics), suggest that Geobatrachus represents an extreme line of specialization among the Eleutherodactylini. The evidence afforded by the cladistic analysis gives no support to regard the Eleutherodactylini are derived from Adenomera or an immediate common ancestor with this genus, and so the acquisition of the direct development among the Leptodactylidae seens to have occurred independently at least in two phyletic lines.Se recopilan las diferentes asignaciones en cuanto a familia que han sido formuladas hasta la fecha para el género Geobatrachus. Mediante la recolección de una serie de topótipos virtuales de este género monotípico, endémico del sector noroeste del macizo de la Sierra Nevada de Santa Marta (Departamento del Magdalena, Colombia), se elaboró una descripción detallada y se procedió a un análisis de 67 características morfológicas, a las cuales se adicionó el numero cromosómico (2n = 20), seleccionadas conforme a las utilizadas por LYNCH y HEYER en sus estudios sobre clasificación y filogenia de los Leptodactylidae, las cuales, después de un examen inicial, se consideraron significativas, a fin de precisar las afinidades de este género, Simultáneamente con la recolección del material se adelantaron observaciones ecológicas que permitieron establecer que su distribución se opera en el área de bosques húmedos, frecuentemente nublados entre los 1.750 y 3.000 metros, y que la especie tolera situaciones paraclimácicas (plantaciones artificiales de Cupressus sp., Pinus sp. y piso cubierto de Pennisetum clandestinum) y que sus hábitos son netamente terrestres y semifosoriales. De los caracteres externos cabe destacar una gran variabilidad cromática individual; el integumento desprovisto de adaptaciones xeromorfas y de complejos glandulares especializados, indica una adaptación hacia un hábitat higrófilo, añadidos a la extremada reducción del primer dedo pedial. Las características esqueléticas determinadas mediante disección y tinción diferencial, muestran un tipo generalizado sin reducción pronunciada de elementos craneales, con presencia de dientes no pedicelados en la arcada maxilar, sin evidencias de adaptaciones fragmóticas ni fosoriales ni ornamentación epicraneal o exóstosis, aparato hioideo generalizado, 8 vértebras presacras sin fusión ni imbricación y con procesos transversos especializados, sacro con diapófisis no dilatadas, astrágalo y calcáneo fusionados, tarsales reducidos a dos.  Es llamativa la presencia de falanges terminales en forma de T, lo cual podría sugerir una pre adaptación hacia un régimen arborícola (o bien la derivación a partir de un antecesor arborícola), Una condición firmisternal plena de la cintura escapular y la notable reducción de la clavícula insinuarían una aparente relación con los Microhylidae.  La musculatura muestra un M. intermandibularis con rafé medial, la inserción del M. adductor mandibularis subexternus de tipo "S", el tendón distal del M. semitendinosus con recorrido ventral a los Mm. graciles; el M. adductor longus con inserción en la rodilla, y la ausencia de una cabeza accesoria o tendón del M. glutaeus magnus (carácter único dentro de los Neobatrachia, pero verosímilmente debido a pérdida secundaria). Una de las características más llamativas (en contra de registros previos de renacuajos atribuidos tentativamente a esta especie) es la confirmación de un desarrollo directo que a priori podría suponerse dada la reducida postura de huevos megalecitales en lugares protegidos y que corresponde a una adaptación hacia ambientes de montaña con escasa oportunidad para un desarrollo larvario acuático.  Los 68 caracteres utilizados fueron codificados numéricamente según sus condiciones primitivas o derivadas, según criterios aceptados en la literatura reciente, tanto para facilitar las comparaciones como para intentar reconstruir la filogenia según principios de taxonomía numérica.  Fácilmente por exclusión se descartó la posibilidad de que Geobatrachus representase conforme a la firmisternia un elemento de los Microhyloidea, y la ausencia de elementos esqueléticos interfalangeales, excluyen cualquier posible afinidad con los Centrolenidae, Hylidae y Pseudidae, aun cuando ninguna característica, salvo la ausencia del tendón accesorio del M. glutaeus magnus (interpretada como pérdida secundaria) excluye la asignación de Geobatrachus a los Bufonoidea. Ninguna característica en particular permite referir el genero a los Myobatrachidae ni a los Heleophrynidae, la ausencia de órgano de Bidder (única característica diagnóstica entre Leptodactylidae y Bufonidae) elimina, aparte de la afinidad mostrada por cladogramas experimentales, la familia Bufonidae de toda su consideración adicional. En cuanto a los Brachycephalidae (sensu McDIARMID 1969), si bien aparentemente derivados de los Leptodactylidae, diferencias tales como la estructura de la cintura escapular  y el desarrollo de osteodermos dorsales los excluyen de consideración.  El análisis combinatorial realizado mediante cladogramas confirma que los Phyllobatinae (= Dendrobatidae) son derivados de los Elosiini, y que los acreditan mejor en el rango de subfamilia dentro de los Leptodactylidae que como familia separada. Aun cuando la firmisternia en particular puede sugerir afinidades con Geobatrachus, ciertamente este género al igual que Rhinoderma no pueden ser asignados a los Phyllobatinae. El análisis de los caracteres de Rhinoderma indica que este género se derivó del antecesor inmediato de los Leptodactylinae o de elementos antiguos de este grupo y que, al igual que en el caso de los Phyllobatinae, bien pueden involucrarse dentro de los Leptodactylidae como una sub familia, a menos que se recurra a dividir dicha familia en varios grupos de jerarquía familiar. No obstante Geobatrachus difiere ampliamente de los Rhinodermatinae y, como lo indican los cladogramas elaborados, muestra su máxima afinidad con los Eleutherodactylini, si bien representa un género bien diferenciado dentro de la tribu quizás como resultado de la evolución in situ de un "stock" antiguo de aislamiento desde el Terciario, que ha sobrevivido en condiciones favorecidas por la ecuabilidad climática, Hipótesis basadas en las evidencias paleogeográficas y paleoclimáticas  disponibles, así como con forme a la posible dicotomía fundamental de los llamados grupos ,.Alpha" y “Beta" de los Eleutherodactylus son discutidas.  Con exclusión de Euparkerella y Holoaden los Eleutherodactylini (cuya diferenciación inicial quizás tuvo lugar en el S. E. de Brasil), los restantes géneros del grupo se hallan estrechamente relacionados entre sí, aun cuando una revisión exhaustiva de Eleutherodactylus que eventualmente puede llegar a subdividir este género, puede llevar a conclusiones que si bien no se aparten substancialmente de las interpretaciones aquí ofrecidas, si conduzcan a una interpretación mejor fundada de la que las hipótesis planteadas o incluso a una reducción del número de géneros actualmente reconocidos.  Tanto la firmisternia como la pérdida del tendón accesorio del M. glutaeus magnus y el número comparativamente reducido de cromosomas con ausencia de acrocéntricos, sugieren que Geobatrachus represente una línea extrema de especialización dentro de los Eleutherodactylini.  La evidencia aportada por el análisis cladístico no favorece considerar a los Eleutherodactylinae como derivados de Adenomera o de un antecesor inmediato común con este género y, por tanto, la adquisición del desarrollo directo dentro de los Leptodactylidae parece haber ocurrido independientemente al menos en dos líneas filéticas.application/pdfspaCaldasiahttp://revistas.unal.edu.co/index.php/cal/article/view/34553Universidad Nacional de Colombia Revistas electrónicas UN CaldasiaCaldasiaCaldasia; Vol. 12, núm. 59 (1979); 383-495 Caldasia; Vol. 12, núm. 59 (1979); 383-495 2357-3759 0366-5232Ardila Robayo, María Cristina (1979) Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura). Caldasia; Vol. 12, núm. 59 (1979); 383-495 Caldasia; Vol. 12, núm. 59 (1979); 383-495 2357-3759 0366-5232 .Status sistematico del género geobatrachus ruthven 1915 (amphibia: anura)Artículo de revistainfo:eu-repo/semantics/articleinfo:eu-repo/semantics/publishedVersionhttp://purl.org/coar/resource_type/c_6501http://purl.org/coar/resource_type/c_2df8fbb1http://purl.org/coar/version/c_970fb48d4fbd8a85Texthttp://purl.org/redcol/resource_type/ARTCiencias NaturalesBiologíaPlantasanimalesHistoria NaturalPaleobotánicaPaleozoologíaOrnitologíaAmphibiaAnuraGénero Geobatrachus RuthvenAdenomeraAmphibiaAnuraGénero Geobatrachus RuthvenAdenomeraORIGINAL34553-133865-1-PB.pdfapplication/pdf44575519https://repositorio.unal.edu.co/bitstream/unal/44306/1/34553-133865-1-PB.pdf980c06e2016bb6f9614907e49787d0b4MD51THUMBNAIL34553-133865-1-PB.pdf.jpg34553-133865-1-PB.pdf.jpgGenerated Thumbnailimage/jpeg7190https://repositorio.unal.edu.co/bitstream/unal/44306/2/34553-133865-1-PB.pdf.jpg64bbac7384b5b46eded058aa49825a24MD52unal/44306oai:repositorio.unal.edu.co:unal/443062024-02-14 23:08:15.224Repositorio Institucional Universidad Nacional de Colombiarepositorio_nal@unal.edu.co